275 Tropical Bryology 8: 275-280, 1993 Notes on the amphipacific relations of Hawaiian Cladoniaceae S. Stenroos Department of Botany, P.O. Box 47, FIN-00014 Helsinki, Finland C.W. Smith Department of Botany of the University of Hawaii at Manoa, Honolulu, Hawaii 96822, USA Abstract. The total number of currently accepted species of Cladoniaceae in the Hawaiian Islands is 22. Several taxonomic problems still exist, however. The effects of isolation are clear among Cladoniaceae. Endemism is high (c. 40%); and, the number of species low. The species must have reached the archipelago via long-distance trans-oceanic dispersal, probably aided by the abundant production of lichen propagules, such as soredia and microsquamules. Although most of the species found in Hawaii are widely distributed, the Hawaiian Cladoniaceae show slight affinities to those of E and SE Asia. Cladonia polyphylla Mont. & v.d. Bosch is an older name for C. fruticulosa Krempelh., and is lectotypified from authentic material. C. leprosula H. Magn. is included in C. ochrochlora Flörke. True oceanic islands, like the Hawaiian Islands, islands (Hawaii, Kahoolawe, Maui, Lanai, Mo- have never had any connection or association lokai, Oahu, Kauai, and Niihau), the largest of with continental land during geographic diversi- which - and still with volcanic activity - being fication that has been going on through geologi- Hawaii (c. 1560 sq km). cal time. These islands are volcanoes arising from the ocean floor. The Hawaiian Islands are Major works on Hawaiian lichens, including shield volcanoes that are formed where tectonic Cladoniaceae, are those of des Abbayes (1947), plates move over a “hot spot” and magma wells Magnusson (1956, who summarizes exhaustive- up to the surface (e.g. Press & Siever 1982, ly the earlier information), Klement (1966) and Fosberg 1991). The estimated age of the Hawai- Smith (1981). Some ecologial investigations ian archipelago is c. 50 million years. Today the including Cladoniaceae have also been reported nearest continent is North America (c. 3800 km (e.g. Kappen & Smith 1980). to the northeast). Japan is c. 5400 km to the west, and the closest oceanic atolls are c. 800 km The main purpose of the present paper is to distant. The archipelago comprises eight major update information on Hawaiian species of the 276 Cladoniaceae, including their taxonomy. In Magnusson 1956), C. pyxidata (L.) Fr., C. addition, we show the importance of long-distan- scabriuscula (Delise) Nyl., C. sphacelata Vainio, ce dispersal that seems to be surprisingly effective and C. subradiata (Vainio) Sandst. (as C. among Cladoniaceae in many areas, especially fimbriata var. balfourii (Crombie) Vainio). Of in the Southern Hemisphere where land is the reindeer lichens, Cladina rangiferina (L.) scattered (see e.g. Galloway 1991, Stenroos 1993). Nyl. and C. mitis (Sandst.) Hustich were definitely excluded from the Hawaiian flora by Ahti (1961). We emphasize that the material used for this study is not complete. Some specimens could not In addition, the following taxa reported from the be determined to any known species, but on the Islands are reduced to synonymy, further redu- other hand we were reluctant to describe new cing the total number of accepted species present: taxa based on sparse material. Others have not Cladonia congregata H. Magn. (= C. didyma been relocated, some destroyed in World War II. (Fée) Vainio, see des Abbayes 1966, Stenroos The results are mainly based on specimens in H, 1986a), C. fauriei des Abb. (= C. kauaiensis HAW and TUR (which include the collection Merrill, see Stenroos 1986a), C. leprosula H. made by L. & Y. Mäkinen in 1966), and on field Magn. (= C. ochrochlora Flörke, T. Ahti pers. work of both authors. Several specimens were comm.), C. subsquamosa (Nyl.) Crombie (= C. analysed by standardized thin-layer chromato- squamosa Hoffm., see Huovinen & Ahti 1988), graphic methods (Culberson 1972, White & James and C. vulcanica Zoll. & Moritzi (= C. didyma 1985). (Fée) Vainio, see Huovinen et al. 1989). C. polyphylla Mont. & v.d. Bosch (as C. pityrea *polyphylla (Mont. & V.D. Bosch) des Abb. in Results and discussion Magnusson 1956) was found here to be conspe- cific with C. fruticulosa Krempelh. However, Altogether 22 species of Cladina and Cladonia although C. polyphylla (lectotype, here selected: are recorded in this study (Table 1). The number Java, Junghuhn, PC-Hue; syntypes, L, PC-Hue) of species reported from the area by earlier is an older name for this taxon, it is not accepted authors is almost twice as high (see Magnusson here because C. fruticulosa shall be proposed for 1956, who recognized 37 species). All previously conservation according to the new Art. 14.1 in recorded species have been assigned to currently the Botanical Code as approved in the XV Inter- extant species in the islands; none are extinct. national Botanical Congress in Yokohama in 1993. The following species reported from Hawaii by earlier authors are excluded from the flora becau- Finally, in addition to Cladina magnussonii (Ahti) se in light of the material examined and the Ahti, which was reported by Ahti (1961), the knowledge of the distribution of the species following taxa are recorded here as additions to elsewhere they are unlikely to occur in the Magnusson’s (1956) list of Cladoniaceae of the Hawaiian Islands, or their presence needs Hawaiian Islands: Cladonia cf. corniculata Ahti confirmation: Cladia aggregata (Sw.) Nyl., & Kashiw., C. merochlorophaea Asah. var. Cladonia chlorophaea (Flörke) Sprengel (as C. merochlorophaea, C. pleurota (Flörke) Schae- pyxidata var. chlorophaea Flörke in Magnusson rer s.lat., and C. poeciloclada des Abb. One 1956), C. coniocraea (Flörke) Sprengel, C. specimen (Kauai, Hanalei, Bowler 1517, H), corallifera (Kunze) Nyl., C. digitata (L.) Hoffm., with farinose soredia, imperfectly scyphose C. erythrosperma Vainio, C. fimbriata (L.) Fr., podetia, and fumarprotocetraric acid, probably C. furcata (Hudson) Schrader, C. gracilis (L.) belongs to an undescribed species (“sp. 1” in Willd., C. granulosa (Vainio) Ahti (as C. Table 1). Specimens (Kauai, Kokee, L. & Y. subsquamosa f. granulosa Vainio in Magnusson Mäkinen 66-2031, TUR; Oahu, Waianae Range, 1956), C. grayi Merr. ex Sandst., C. macilenta L. & Y. Mäkinen 66-1627, TUR; Hawaii, Hoffm., C. mexicana Vainio, C. phyllophora Volcanoes Nat. Park, Stenroos 3918, H) Hoffm. (as C. degenerans (Flörke) Sprengel in determined here as “sp. 2” are related to C. 277 Table 1. List of Hawaiian Cladoniaceae and their major secondary substances. BAR=barbatic acid (incl. 4-O-demethylbarbatic acid), DID=didymic acid, FUM-complex=fumarprotocetraric and protocetraric acids and Cph-2 (Cph- 1 present or absent), HSEK=homosekikaic acid, IUSN=isousnic acid, MCRY=4'-O- methylcryptochlorophaeic acid, MER=merochlorophaeic acid, PER=perlatolic acid, PSO=psoromic acid, SQU=squamatic acid, THA=thamnolic acid, USN=usnic acid. ———————————————————————————————————— —————— Species Major secondary substances ———————————————————————————————————— —————— Cladina leiodea (H. Magn.) Ahti USN, FUM-complex magnussonii (Ahti) Ahti ATR, FUM-complex skottsbergii (H. Magn.) Follm. USN, PER f. fuscescens (Ahti) Ahti PER Cladonia angustata Nyl. USN, BAR cf. cenotea (Ach.) Schaerer SQU cf. corniculata Ahti & Kashiw. FUM-complex didyma (Fée) Vainio THA, DID or BAR farinacea (Vainio) A. Evans FUM-complex fruticulosa Krempelh. PSO or FUM-complex kauaiensis G.K. Merrill ex H.Magn. & Zahlbr. USN, THA mauiensis H. Magn. BAR, DID merochlorophaea Asah. var. merochlorophaea MCRY, MER, FUM-complex oceanica (Vainio) Zahlbr. USN, BAR, DID ochrochlora Flörke FUM-complex pleurota (Flörke) Schaerer USN, IUSN, ZEO poeciloclada des Abb. FUM-complex ramulosa (With.) Laundon FUM-complex, HSEK, ATR (incl. C. adspersa Mont. & Bosch) solitaria H. Magn. USN squamosa Hoffm. SQU subsquamosa Krempelh. FUM-complex ramulosa and C. phyllopoda (Vainio) Stenroos as well. However, this complex still needs a (Stenroos 1988) but differ from them by their thorough revision. For some of the more poorly very slender, more branched, granulose, and known taxa of those accepted here, the reader is partly escyphose podetia. Specimens generally referred to some previous papers: Cladonia referred to as C. scabriuscula or C. farinacea are angustata Nyl. (Stenroos 1986a), C. fruticulosa here included in C. farinacea s.lat. The present Krempelh. (as C. decipiens des Abb. in material is often richly squamulose (in contrast Magnusson 1956, see Stenroos 1988), C. to C. farinacea s.str. in South America) and the mauiensis H. Magn. (Stenroos 1986a), C. podetial squamules and parts of podetial surface oceanica (Vainio) Zahlbr. (Stenroos 1986a), and are farinose-sorediate (contrary to C. scabriuscula C. solitaria H. Magn. (Stenroos 1986b). s.str.). Similar material is found in Eastern Asia 278 The effects of prolonged isolation of the Hawai- have weak affinities with Central and South ian Islands are clear in taxa of the Cladoniaceae. America (Smith 1993, 1994). The degree of endemism in lichens is high, viz. c. 40% (parmelioid lichens 8%, alectorioid li- Successful dispersal is not only a problem of how chens 0%, umbilicarioid lichens 0%, lichens to reach an area but also of establishment once total 38% (see Magnusson 1956), mosses 46% there. Asexual propagules, produced abundantly (see Hoe 1979), ferns & fern allies 70% (C. H. in most Cladoniaceae, have both bionts present. Lamoureux, pers. comm.), and angiosperms Yet most species are slow in their development 95%), and